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The Species of Wurmbea

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J. Adelaide Bot. Gard. 16: 33-53 (1995) THE SPECIES OFWURMBEA(LILIACEAE) IN SOUTH AUSTRALIA Robert J. Bates Cl- State Herbarium, Botanic Gardens, North Terrace, Adelaide, South Australia 5000 Abstract Nine species of Wunnbea Thunb. are recognised in South Australia. W. biglandulosa (R. Br.)Macfarlane, W. deserticola Macfarlane and W. sinora Macfarlane are recorded for the first time; Wurmbea biglandulosa ssp. flindersica, W. centralis ssp. australis, W. decumbens, W. dioica ssp. citrina, W. dioica ssp. lacunaria, W. latifolia ssp. vanessae and W. stellata are described. A key, together with notes on each species is provided. Macfarlane (1980) revised the genus for Australia. He placed Anguillaria R. Br. under Wurmbea and recognised W. dioica (R. Br.)F. Muell., W. centralis Macfarlane, W. latifolia Macfarlane and W. uniflora (R. Br.)Macfarlane as occurring in South Australia. Before this only one species, W. dioica (as Anguillaria dioica) was listed for South Australia (J.M. Black 1922, 1943). Macfarlane stated that he had seen no live material of South Australian species. The present author has made extensive field studies of taxa discussed in this paper, has cultivated most and studied herbarium material. Several trips have been made to other states to allow further comparisons to be made. For information on the nomenclatural history, general morphology, biology and ecology of Wurmbea see Macfarlane 1980. Key to the South Australian species of Wurmbea 1 Lower leaves paired (almost opposite), basal, of same shape and size 2 1: Lower leaves well separated, often of different shape and size 4 2 Leaves with serrate margins, flowers unisexual, nectaries 1 per tepal, a single band of colour.... 6. W. latifolia 2:Leaves without serrate margins, flowers hermaphroditic, nectaries 2 per tepal 3 3 Lower leaves narrow-linear, decumbent, flower 1, < 7 mm across, capsule elongate on a decumbent stem 3. W. decumbens 3:Lower leaves broadly linear-lanceolate, flowers several, > 7 mm across, capsule ovoid on an erect stem 2b. W. centralis ssp. australis 4 Nectary 1 per tepal, forming a single band of colour, flowers unisexual 5. W. dioica 4:Nectaries 2 per tepal, flowers hermaphroditic 5 5 Flower single, tepals <3 mm broad 6 5:Flowers several, tepals > 3 mm broad 8 6 Flower < 10 mm across, tepals clasping filament (at least in living plant), outer margin of nectary winged 7 6:Flower > 10 mm across, tepals not clasping filament, outer margin of nectary not winged 8. W. stellata 7 Basal leaf linearLlanceolate, receptacle thickened, anthers yellow, spring flowered 9. W. un flora 7:Basal leaf filiform, receptacle not thickened, anthers purple, winter flowered 7. W. sinora 8 Tepals connate, forming a cup-shaped tube up to 1/3 of length, nectaries small, elliptical with margins elevated all round, styles connate 4. W. deserticola 8:Tepals not connate, not forming a cup-shaped tube, nectaries large, not elliptical, without elevated margins all round, styles not connate 9 9 Flowers moderately crowded, nectaries semi-oval, desert plants 2a. W. centralis ssp. centralis 9:Flowers well spaced, nectaries like broad ledges, southern plants 1. W. biglandidosa 33 R.J. Bates J. Adelaide Bot. Gard.16 (1995) det daAar-sraa Fig. 1. Habit of dioica ssp. citrinaWurmbea species (natura]size). A, W. biglandulosa (1 W. Rogers 464);C, W decumbens (R. ssp. Andersica (R. Bates Bates 10483); E, W Bates 25630); D, W. 10264); B, W latifolia ssp. latifolia,male (R Copley 5213); dioica ssp. lacunaria, 25606); G, W stellata(R. Bates 19760). F, W. latifolia male (R. ssp. vanessae, male (RBates 34 J. Adelaide Bot. Gard. 16 (1995) Wurmbea in South Australia Fig. 2. AI Flower. A, Wurmbea biglandulosa ssp. flindersica (R. Bates 10264); B, W. cenn-alis ssp. australis (R. Bates 40825); C, W. dec-umbens (R. Bates 28725); D, W. dioica ssp. citrina (R. W. Rogers 464); E, W. dioica ssp. lacunaria (R. Bates 10483); F, G, W. latifolia ssp. vanessae (R. Bates 25606); F, male flower; G, female flower; H, W. sinora (R Bates 26263);1, W. stellata (R. Bates 19760). JM, Fruits. J, W. decumbens (R. Bates 25672); K, W. dioica ssp. citrina (R. Bates 20788); L, W. latifolia ssp. latifolia (B. Copley 5213); M, W. latifolia ssp. vanessae (R. Bates 32462). 35 R.J. Bates J. Adelaide Bot. Gard. 16 (1995) 1. Wurmbea biglandulosa (R. Br.)Macfarlane, Brunonia 3 (1980) 191; FI. Austr. 45 (1987) 400. Anguillaria biglandulosaR. Br., Prodr. 273 (1810). Anguillaria australisF. Muell., Fragm. 7 (1870) 74, p.p.,nom. illeg. Melanthium browniiSchltdl., Linnaea 1 (1826) 86, p.p.,nom. Meg. W. biglandulosa has not previously been recorded for South Australia despite having been collected here over one hundred years ago and being locally common. The type form does not appear to occur here, and the South Australian plants are described as a separate subspecies. Key to subspecies 1 Uppermost leaf long, acuminate, flowers white, thin textured, nectaries clasping filaments (in living plants) la. ssp.biglandulosa 1:Uppermost leaf short, acute, flowers pink (at least outside) thick textured, nectaries not clasping filaments lb.ssp.flindersica lb. ssp. flindersica R. Bates, ssp. nov. (Figs 1A, 2A) A ssp. biglandulosahabitu robusto, tepalis longioribus crassis roseis (non albis),nectariisfilo non amplectantibus et foliis supremis brevior differt. Type: On burnt ground, roadside opposite Army Camp at Hughes Gap (NL), 10.viii.1992, A.G. Spooner 13351 (holo.: AD; iso.: AD, MEL, PERTH). Plants moderately large, 5-30 cm tall. Corm ellipsoidal, 1.5-2 x 0.8 cm, 1-4 cm below ground.Lecrves3,well separated;lowest one basal,hardly dilated at base, linear, 5-15 cm long, 2 4 mm diam. at middle; middle leaf shorter dilated at base, upper portion long and tapering, filiform; uppermost leaf markedly dilated at base, with a short acuminate apex, attached well below the inflorescence. Flowers 2-6, all hermaphrodite. Perianth deep-pink outside,paler pink or whiteinside. Tepals 6, up to 12 mm long, shortly connateatbaseorfree,segments narrow below nectaries, broader and spreadingbeyond,narrowelliptic. Nectaries 2 per tepal, brilliant carmine, situated 1/3 from base, in shallow pits, separated at the centre and reaching to margins, not clasping filaments. Stamens two thirds length of tepals, Map. 1. Distribution of W.biglandulosa ssp.flindersica & W. sinora 0in South Australia. 36 J. Adelaide Bot. Gard. 16 (1995) Wurmbea in South Australia filaments adnate to perianth only near base, not swollen. Anthers oblong, 2 mm long, versatile, attached at middle, deep purple-red. Ovary oblong, carpels sharply delimited from the free styles. Capsules dehiscing loculicidally. Seeds dark brown, 2 mm diam. Distribution and ecology (Map 1) Endemic to the southern Flinders Ranges from Gladstone north to Quorn, a distance of about 100 lun. Common in open woodland dominated by Eucalyptus cladocalyx and Callitris glaucophylla usually on rocky slopes in fertile soils; in places forming a dominant ground cover after fire. Flowers: August to October; perfumesweet, honey-like. Distinguishing features Easily recognised by the 2 large pink nectaries on each tepal. It can be separated from ssp. biglandulosa (which is confined to the Eastern States) by its pink rather than white flowers, the thick textured tepals with brilliant pink nectaries which do not clasp the filaments, and the shorter top-leaf. The 2 subspecies are separated geographically by over 500 km. Variation: A rather constant taxon, varying only in number and size of flowers. In good seasons or after fires plants produce more and larger flowers. Sympatric species W. biglandulosa ssp. flindersica is often found with W. centralis but flowers 1-4 months later. Near Wirrabara it flowers together with W. dioica ssp. dioica in August and north of Melrose it occurs with W. latifolia ssp. latifolia in Eucalyptus camaldulensis woodland. Hybrids have not been recorded. Etymology: As subspecies flindersica is only found in the Flinders Ranges the epithet is appropriate. Conservation status: Common and well conserved throughout its range. Notes Macfarlane (1980, 84) did not record W. biglandulosa from South Australia. He identified some collections, i.e. 11.M. Cooper Melrose' as W. centralis and his reference to pink flowered forms of W. dioica from South Australia may have been due to notes on collections of W. biglandulosa. Both flies and native bees have been observed pollinating W. biglandulosa near Telowie Gorge. Selected specimens seen (from c. 50 seen at AD) SOUTH AUSTRALIA. FLINDERS RANGES: Alligator Gorge, 12.ix.1987, R. Bates 10264; Melrose, ix.1960, 11.M. Cooper s.n.; Mt Remarkable, -.ix.1974, 11.M. Cooper s.n.; HOROCICS Pass, 29.xii.1922, T.G.B. Osborne s.n. (this bears the collector's label...`Anguillaria dioica var. hermaphrodita'); Mambray Creek, -.x.1960, J. Shillabeer s.n.; On ridge N side of Mambray Creek, 5.ix.1974, D.JE. Whibley 4270; N slopes of Mt Brown, 20.x.1958, P.G. Wilson 623. NORTHERN LOFTY: Telowie Gorge, 25.ix.1989, R. Bates 20543; Near the Bluff E of Port Pire, 1.x.1978, A.G. Spooner 6019; 10 km N of Gladstone in Explosives Reserve, 7.ix.1991, R. Bates 24865. 2. Wurmbea centralis Macfarlane, Brunonia 3 (1980) 188; in Jessop, J.P. (ed.), Fl. Central Austr. 422 (1985); in Jessop, J.P. & Toelken, H.R. (eds), FI. S. Austr. 4 (1986) 1772; Fl. Austr. 45 (1987) 395. 37 R.J. Bates J. Adelaide Bot. Gard. 16 (1995) According to Macfarlane (1980) the type location of this species (Mt Olga) `... is far from the nearest South Australian locality.' and he predicted that it would occur in the Musgrave and Everard Ranges of South Australia. Macfarlane's concept of W. centralis (1980, 1987) was very broad, he had seen no live material and included under W. centralis collections from as far afield as Kangaroo Island and the southern Flinders Ranges.
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    MADS-Box Protein Complex Vvag2, Vvsep3 and Vvagl11 Regulates the Formation of Ovules in Vitis Vinifera L

    G C A T T A C G G C A T genes Article MADS-Box Protein Complex VvAG2, VvSEP3 and VvAGL11 Regulates the Formation of Ovules in Vitis vinifera L. cv. ‘Xiangfei’ Yan Wang, Zhenhua Liu, Jiang Wu, Liang Hong, Jinjun Liang, Yangmei Ren, Pingyin Guan and Jianfang Hu * College of Horticulture, China Agricultural University, Beijing 100193, China; [email protected] (Y.W.); [email protected] (Z.L.); [email protected] (J.W.); [email protected] (L.H.); [email protected] (J.L.); [email protected] (Y.R.); [email protected] (P.G.) * Correspondence: [email protected]; Tel.: +86-010-6273-2488 Abstract: The phenomenon of multi-carpel and multi-ovule exists in the grapevine cultivar ‘Xiangfei’, but the mechanism of ovule formation is seldom reported. In this study, we observed the ovule formation process by using ‘Xiangfei’ grapes. The role of the VvAG2 (VvAGAMOUS) gene in ovule formation was identified, and we explored the relationship between VvAG2, VvSEP3(VvMADS4) and VvAGL11(VvMADS5) proteins. The results showed that the ovule primordium appeared when the inflorescence length of ‘Xiangfei’ grapes were 4–5 cm long; the relative expression levels of VvAG2, VvAGL11 and VvSEP3 genes were higher during ovule formation, and the expression levels of VvAG2 gene was the highest. Transgenic tomato (Solanum lycopersicum) plants expressing VvAG2 produced higher numbers of ovules and carpels than the wild type. Moreover, yeast two-hybrid and yeast three-hybrid experiments demonstrated that VvSEP3 acts as a bridge and interacts with VvAG2 and VvAGL11 proteins, respectively. Meanwhile, a homodimer can be formed between Citation: Wang, Y.; Liu, Z.; Wu, J.; VvSEP3 and VvSEP3, but there was no interaction between VvAG2 and VvAGL11.
  • Flowers and Flower Terminology Biology 205, Spring 2006

    Flowers and Flower Terminology Biology 205, Spring 2006

    Flowers and Flower Terminology Biology 205, Spring 2006 A flower is a terminal cluster of sporophylls, and, technically, a strobilus. A flower is composed of 4 whorls of sporophylls 2 sterile (sepals and petals) 2 fertile (stamens and carpels) Collective Terms All sepals = calyx All petals = corolla All sepals and petals together = perianth If you can’t tell between sepals and petals, use the term tepals (e.g., Magnolia flowers) All stamens = androecium All carpels = gynoecium Stamens are made up of the filament (sporophyll remnant) and anthers (containing pollen) Carpels are made up of the stigma (receptive surface), style (neck) and ovary (containing ovules) The placenta is the tissue in the ovary that takes nutrition to ovules/developing embryo Placentation is the arrangement of ovules along the placental tissue A primitive stamen with a pronounced filament is called a laminar stamen (not found in more advanced flowers) A primitive carpel that is not fully fused is called a conduplicate megasporophyll (not found in more advanced flowers) The two major trends of advanced flowers are reduction and fusion Reduction: in the # of parts or in the loss of whole parts (whorls) Fusion: of like parts (connation) or unlike parts (adnation) Fusion of calyx and corolla is usually into a tube Asepalous (free sepals) synsepalous (fused sepals) Apetalous (free petals) synpetalous (fused petals) (the prefix A- means free parts, Syn- means fusion of like parts) Stamens can be fused together by their filaments (as in many legumes) or by their anthers